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Home , Defaunation, Emas National Park

Parks of the Pleistocene: recreating the and the with megafauna

in the archeological record) and the of great Parks of the Pleistocene: birds and the world over. An important fact to observe is that, although the climatic changes at recreating the Cerrado the end of Pleistocene happened at the same time all over the world, the extinction of the megafauna began and the Pantanal with 40,000 years ago in and Australia, 12,000 years megafauna1 ago in America, and less than 1,000 years ago in Madagascar and New Zealand (Martin, 1995, Flannery, Mauro Galetti, PhD2,3 Phenology and Seed Dispersion 1995). Group, Department of Ecology (UNESP-Rio Claro) However, other researchers suggest that climate changes alone could have exterminated the megafauna In the past few years, several (ex. Cartelle, 1999; Ficcarelli et al, 2003). With the researchers have proposed increase of rain and temperature, the open savannas the introduction of large would have been reduced in size, detrimentally affecting predators and herbivores to facilitate the reconstruction the expansion of closed humid forests, causing the of natural . The introduction of wolves, elimination of the megafauna due to a lack of favorable bears, pumas and even nonnative species such as environment. These researchers don’t accept the and camels, has been suggested by hypothesis that primitive man caused the extinction of researchers to reestablish the ecological balance in the megafauna, alleging that there isn’t a correlation several North American ecosystems, from forests to with megafauna bones. However, areas with a human- deserts (Martin & Burney, 2000). Even the well- megafauna interaction, though not abundant, exist, equipped North American national parks failed to although this evidence is discredited (see Fiedel & protect the entire diversity of great mammals such as Haynes, 2004). Beyond this, in many places the bears, coyotes and wolves (Newmark, 1987). megafauna is not restricted to open spaces; elephants, Predators at the top of the food chain are considered and other ungulates can be found in to be “keys” to the maintenance of the , closed forests. because they regulate the population of the herbivores. It is most probable that both and climatic Without these predators, there is a disproportionate changes played important roles in the extinction of the increase in the populations of herbivores and South American megafauna. Owen-Smith (1992) mesopredators (foxes, raccoons, and squirrels), which proposes that the extinction of the Pleistocene overtax the population of plants and small animals which megafauna involved climatic factors as much as are their foods (Terborgh, 1992; Crooks & Soulé, anthropic ones. Widespread hunting by the Paleo- 1999). Indians would have caused an accentuated decline in Many years before the arrival of the first humans to the megaherbivore population (as happens today in America, the North American savannas held at least several indigenous reservations and protected areas 41 species of great mammals, including wild horses, with a great deal of hunting). The changes in the bisons, camels, giant armadillos, mammoths and giant climate, however, would have fragmented and restricted sloths (Anderson, 1995), while in that the distribution of those megaherbivores to places of diversity was much larger (Cartelle, 1999; Fariña et al., low nutritional value, leaving them more vulnerable to 1998). local extinction as much to human, as to stochastic factors. These mega-mammals played an extremely important role in the structure of vegetation communities and In addition to this, extinct mammals possess low the effects of their premature extinction are still poorly reproductive rates, except for nocturnal or arboreal understood (Janzen & Martin, 1982; Janzen, 1986). species (Johnson, 2002). The mammals and birds Some researchers suggest that there is sufficient dependent on the megafauna (such as sabre-tooth evidence to demonstrate that primitive man played a tigers and condors) would have been wiped out as a primary role in the extinction of mammoths, giant sloths, result of the extinction of their prey. camels, llamas, glyptodonts, horses and other species Actually, we still know little about what factors of large mammals, called “megafauna” (Martin, 1995; caused the extinction of the megafauna, but, perhaps Haynes, 2001). A strong correlation exists between the most pertinent question is what are the colonization and growth of human populations (visible effects of the extinction of the megafauna on neotropic savanna ecosystems. 1 Sent originally in Portuguese 2 [email protected] With the rapid elimination of South American megafauna 3This work relied upon the support of Carina Denny, Paulo Guimarães Jr, Richard Fariña and Fábio Olmos and is dedicated to Megafauna Team. Mauro Galetti (some authors suggest between eight and three receives CNPq scholarship funding and support from FAPESP and IFS. thousand years ago – De Vivo & Carmignotto, 2004),

Natureza & Conservação . vol 2 . n1 . april 2004 - pp. 93-100 95 Point of View Mauro Galetti

the absence of large herbivores increased the Based upon what we know about South American of some plants, resulting in an accumulation fossils, and knowing the ecological role of the of dry vegetation biomass susceptible to fire. With the megaherbivores in Asia and Africa, certainly all the extinction of easy and naïve prey, such as the large ecosystems, which we know today, particularly the megaherbivores, primitive man intensified the use of Cerrado and the Pantanal, are the result of a massive fire to facilitate the hunting of other mammals, initiating defaunation of great mammals and environments highly the alteration of the savanas (or, that is, the Cerrado) altered by primitive man. as we know them today. It is well known that all of the groups indigenous to the savannas use fire to hunt What does the extinction of the megafauna thousands deer and giant anteaters (Prada, 2001). This type of of years ago have to do with the conservation of today’s hunting has such an impact that some species of large savanna ecosystems, like the Cerrado and the Panta- mammals are already extinct in indigenous reserves nal? The rapid removal of a diverse and abundant (Leeuwenberg, 1997). megafauna, responsible for a good deal of the wealth of species and certainly for most of the My generation grew up greatly influenced by “Animal biomass of those ecosystems, is reflected until today Planet” and other nature programs, all, almost without in the ecological processes of the neotropic savannas. exception, about African savannas. I grew up thinking A comparison between the Emas (Par- Africa was the continent of mammals, while South que Nacional de Emas) in Goiás state with the Kruger America was the continent of birds. The great African National Park in South Africa, or with other African parks, like Serengeti, Okavango, Ngorongoro and parks, can elucidate and open up new horizons in our Kruger, with their herds of zebras, elephants, gnus understanding of our ecosystems. and so many other mammals, were the image of wild nature. Emas National Park is a savanna, 132 Thousand hec- tares (326 acres) in size, which has a significant diversity Large mammals have always attracted, and still attract, of large mammals including the pampas deer, the tapir, the interest of most people. One proof of this is the and the (Silveira et al., 1999). Emas is considered large number of people that visit the or to be the “Brazilian Serengeti” due to its similarity to enclosures at zoos, compared to the the African park. numbers that visit the manned wolves or the capybaras. South American biologists have always Any visitor will observe small groups of pampas deer been slightly jealous of African biologists, because of (Ozotocerus bezoarticus) feeding on the vegetation the lack of large animals in the Neotropics. Few people (Rodrigues, 2003). It is estimated that the park shelters are aware, however, that the population of large South about 1,300 pampas deer, that is, a biomass of 35 kg/ American mammals was, at one point, much richer than km2 (199 lbs/sq. mi.) (Rodrigues, 2003). If we include that found today in Africa. tapirs, white-lipped , red brockett deer and brown brockett deer, the biomass of “large” mammals If we could go back in time at least ten thousand years, doesn’t exceed 100 kg/km2 (570 lbs./sq. mi.) (F. H. G. to the end of the Pleistocene, the savannas of South Rodrigues, pers. comm.). In the African parks, on the America (such as the Cerrado and the Pantanal) were other hand, the biomass of large herbivores can vary more spectacular than the savannas of Africa. Whereas from 5,000 to 22,500 kg/km2 (28,500 to 128,250 lbs./ in Africa there exist only five mammals of more than a sq. mi.), where gnus, zebras, elephants, rhinoceroses, ton (elephant, two species of rhinoceroses, hippo- impalas and other mammals are easily seen (Caro, potamus and the male giraffe), in South America during 1999). the Pleistocene there were more than 38 genera above 100 kilos (220 lbs.), and from ten to twelve species In the Pantanal, known as the place with the largest above a ton, in one single place (Fariña et al., 1998). abundance of mammals in the neotropics, the biomass of the wild herbivores (like pampas deer, deer and Herds of horses (Equus and Hippidion), giant sloths capybaras) doesn’t exceed 1,000 kg/km2 (5,700 lbs./ weighing up to five tons (Eremotherium), mastodon, sq. mi.) (Tomás et al., 2001; Mauro et al., 1998; Galetti which were similar to four ton elephants (Stegomastodon et al., data not published), while the biomass of and Haplomastodon), Xenorhynoterium, which looked nonnative animals, like cattle, feral hogs and buffaloes like one ton camels, the Toxodon, similar in size and can reach more than 5,000 kg/km2 (28,500 lbs./sq. features to the hippopotamus, giant armadillos of up to mi.) (not counting the horses) (Mourão et al., 2002). two tons (Glyptodon), and capybaras weighing 150 This high biomass of nonnative megafauna is sustained kilos (330 lbs. - Neochoerus) wandered our savannas by natural pastures, that certainly should have held and the Pantanal (as well as Caatinga4 and southern large herbivores in the Pleistocene. fields) (Fariña et al., 1998; Cartelle, 1999). It is estimated that the North American savannas before the beginning of the Holocene sustained about 4 Type of vegetation common to the arid regions of the 9,000 kg/km2 (51,300 lbs./sq. mi.) of herbivores, such

Point of View 96 Natureza & Conservação . vol 2 . n1 . april 2004 - pp. 93-100 Parks of the Pleistocene: recreating the Cerrado and the Pantanal with megafauna

as mammoth, horses, bison and other large mammals (megafauna recently introduced) are the only species (Martin, 1995). In the area of the pampas, it is that disperse large fruit in the Pantanal. estimated that there was an historic biomass of mammals of 15,500 kg/km2 (88,350 lbs./sq. mi.), of Currently, one of the largest problems at the Emas which 11,000 kg/km2 (62,700 lbs./sq. mi.) represented National Park, as with many protected areas in the just the megaherbivores (larger than one ton - Fariña, Cerrado and the Pantanal, is fire, caused by lightning 1996). or by man. The Pantanal, with an area of 140,000 km2 (54,040 It is estimated that most of the fires in Emas in the last sq. mi.), consists of 31% natural pasture (around ten years, were natural fires caused by lightning 43,400 km2 (16,752 sq. mi.)) (Silva et al., 2000). Using (Clayish, 2000). In 1994, the whole National Park data regarding the support capacity the natural burned, causing the death of about 330 giant anteaters pastures of the Pantanal offer to cattle, that is, how (Silveira et al., 1999). The park is, in its largest expanse, many kilos of food each square kilometer (of pasture) an open savanna covered by arrow grass (Clayish, can offer to the herbivore fauna, the results indicate 2000). that those natural pastures could sustain about The wild mammals currently at Emas do not consume 10,000 kg/km2 (57,000 lbs./sq. mi.) of megaherbivores this grass (Rodrigues & Monteiro-Filho, 1999), which (if we stipulate a density of 29 cows/km2 (75 cows/sq. accumulates a high biomass and in the dry period mi.) and each cow weighs 350 kg (771 lbs.)) (Santos becomes an ideal fuel for large burnings. et al., 2002). In decade of the 70’s, when the Emas National Park This value is similar to some of the productive African was not fenced and was regularly invaded by cattle savannas, but due to the seasonal and environmental and horses from neighboring farms, neither fire nor differences of the Pantanal, it is probable that the arrow grass were common. (Silveira et al., 1999). ecosystem supports a larger biomass than the African ecosystem (see Fariña, 1996, for the southern fields of The removal of the cattle caused large problems for the Paraguay). park: a higher incidence of fire and the invasion of Even if we take into account that ten thousand years nonnative plants. This isn’t to say that cattle are the ago the Pantanal was drier and colder, very different solution to fires in the Cerrado, but it does raise the from today, and that there are large differences in question of the importance of nonnative species in the productivity within the Pantanal due to flooding, the conservation of local . productivity of this ecosystem could support a high This same effect is found in temperate savannas, where megaherbivore biomass. the removal of large herbivores resulted in an There are several other indicators, which suggest that accumulation of combustible material and a reduction in the Cerrado and the Pantanal are phytoformations that the diversity of pastures (Hartnett et al., 1996, Kramer et co-developed with large mammals, today extinct. In al., 2003). Today, the conservation projects for prairies the Cerrado as well as in the Pantanal, the plants and areas of open vegetation use a combination of demonstrate several adaptations against the controlled burning and the introduction of bison herds herbivorous practices of large mammals, such as thorns (and even cattle) to maintain the mosaic of and on their leaves and trunks (Janzen & Martin, 1982; to avoid the loss of species and harmful fires. Galetti, data not published). There is still a large controversy whether fire is a “natu- In addition to this, many species have fruit which are ral” or simply an anthropic effect in the Cerrado and in too big and well-protected. Fruits like suari nut, other ecosystems (Caldararo, 2002; Svenning, 2002). bocaiúva, indaiá and several other species are too big Traces suggest that fire has been occurring in the Cer- to be consumed and dispersed by the current fauna rado for more than 30,000 years, but about 8,000 (Guimarães & Galetti, 2001). Since 2002, Prof. Marco years ago (when primitive man already occupied the Pizo, of the Department of Botany of UNESP; Camila Cerrado) fire began to occur with much greater Donatti, of the Biology of Conservation Institute (Ins- frequency, indicating an increase in the human tituto de Biologia da Conservação) and I are studying population of the area (Ledru, 2002). the morphology of fruits in the Pantanal, and comparing We know that the Xavantes Indians and other it to our Atlantic database (Galetti et al., ethnicities that live in the Cerrado, as well as the first 2002). people that occupied that area, constantly use fire to It is well-known that the fruits of the Pantanal and hunt, for war and to renew the crops (Prada, 2001). Cerrado are larger and almost always yellow, brown or Whether fire is natural or not, the fact is that the Cerra- orange; colors typical of dispersion. The tapir do, currently, without megaherbivores, accumulates a (the last wild representative of the frugivorous- high biomass of fuel annually, which burns easily and herbivore megafauna), as well as cattle and feral hogs quickly.

Natureza & Conservação . vol 2 . n1 . april 2004 - pp. 93-100 97 Point of View Mauro Galetti

Several studies in Asia, Africa and North America have megaherbivores. In the African savannas it has been demonstrated that large herbivores exercise a great shown that the megaherbivores contribute greatly to influence over the vegetation, regulating the abundance the quality of the soil (McNaughton, 1976). The of competitively superior species, and thus, in low nutrients in the feces and urine of these mega- densities, can increase the local diversity (Harrison et herbivores are immediately available to the plants, al. 2003). In the African savannas the great herds of accelerating the nutrient cycling of the ecosystem. herbivores, such as elephants, buffaloes, impalas, gnus and zebras consume a large portion of the biomass of Beyond this, it is quite reasonable to assume that great grasses, leaving little fuel for possible natural fires. megaherbivore herds must have migrated across the Cerrado and the Pantanal seeking for more adequate Anyone can foresee what would happen if we removed pastures, as happens today with cattle. During the all of the megaherbivores from the Serengeti or from flood periods in the Pantanal these megaherbivores the Kruger. There would be a brutal increase in the probably migrated to higher areas, such as the Cerra- vegetation biomass, the vegetation would do. In the Serengeti it is estimated that a million gnus, be simplified, resulting in a few species becoming 300 thousand zebras and another 300 thousand dominant and, even with a subsequent increase in impalas migrate to the African savanna, feeding about herbivore insects (such as termites and leaf-cutter three thousand lions (Wolanski et al., 1999). ants) the area would have high propensity for fire. Migrations of large herbivores, in reality, happen on all The megaherbivores’ impact is sufficient to shape the continents, from the frozen tundras of the Tibetan structure of an entire ecosystem. The different divisions plateau to the African savannas and, in the past, to the of the physiognomies in the African savannas are based neotropical savannas (Berger, 2004). Currently, the as much on the fertility of the soil as on the impact of only species of large mammal that migrates is the feral fauna (mainly elephants) and on a regimen of burnings hog (Peres, 1996). The interruption of this system of (van Langevelde et al., 2003). In , the definitions migration certainly must have affected the genetic flux of the types of savanna (such as campo–cerrado - via seeds of several plants in the Cerrado and in the scattered in dense grass, cerradão - dense Pantanal. It is not uncommon for geneticists to savanna woodlands, campo-sujo - grassy scrublands, conclude that some species of trees in the Cerrado etc.) take into account just abiotic factors (such as possess low interpopulational genetic variability soil fertility, pluviosity and the frequency of burnings), probably due to a reduction in the genetic flux via seeds, and neglect the effect that the megaherbivores would as already demonstrated by the pequi (Caryocar have on every type of vegetation. brasiliensis) (Collevatti et al., 2003).

One of the ways for us to evaluate the past impact of Therefore, programs for the conservation of the Cerra- large mammals in the Cerrado and in the Pantanal is to do and the Pantanal should take into account the compare it to the modern analogy of the African conditions of these ecosystems before the arrival of savannas, even though the African savannas are much the first humans. What do we want to preserve? The poorer in megafauna than the South American Cerrado as it is today, with an incomplete complement savannas of the Pleistocene. An African elephant of fauna that is difficult to manage and full of vague weighing three tons can consume 150 kg (331 lb.) of niches, which is a direct reflection of the great wave of vegetation a day, returning 135 kg (298 lb.) of fertilizer extinction at the beginning of Holocene, or try to recreate to the environment (Owen-Smith, 1992). an ecosystem that co-developed over millions of years possessing a great diversity and biomass of mega- At least seventy species of fruit are dispersed by forest herbivores? It is well-known that large herbivores have elephants, and some species show a very low a fundamental role in the structuring of the savannas replenishment rate in the absence of elephants (Ale- all over the world and conservationist biologists and xandre, 1978; Nchanji & Plumptre, 2003). The decision-makers cannot ignore this fact. mastodons, members of the family of neotropic elephants which lived in America about ten million years Currently, there is a group of researchers that propose ago and succumbed to the human invasion only 13,000 the use of the fire to manage the Cerrado (see Ramos- years ago, must have had a substantial impact on the Neto & Pivello, 2000). I believe that the use of fire as a vegetation of the Cerrado and the Pantanal just as means to manage the Cerrado can cause greater they had in the North American prairies (Haynes, 2002). damage to the planet, releasing tons of carbon How much could a mastodon weighing four tons con- monoxide into the atmosphere, thus worsening the sume, or even a giant sloth, which could weigh up to greenhouse effect. A large part of the carbon monoxide five tons, and what are the implications of the biomass released into the atmosphere comes from burnings in cycle caused by these megaherbivores? the savannas. An alternative could be the management of the Cerrado and the Pantanal through the re- Therefore, the low fertility of the soil in the savannas introduction of megaherbivores. As giant sloths, could be partially explained by the absence of toxodons, macrauchenias and mastodons don’t exist

Point of View 98 Natureza & Conservação . vol 2 . n1 . april 2004 - pp. 93-100 Parks of the Pleistocene: recreating the Cerrado and the Pantanal with megafauna

anymore, we would have to turn to their closest relatives, In addition, large predators, such as lions, would be or ecological equivalents, even if they are considered excluded from these parks and the population of to be “exotic.” herbivores would be controlled by the park manage- ment, as occurs in African parks. In a short time, the Certainly the introduction of horses, impalas, elephants “Pleistocene Parks” could attract as many tourists as and other megaherbivores into our savannas will shock the African parks, help to reduce the release of carbon many conservationists, but I believe that controlled monoxide caused by burnings, and produce valuable experiments in confined areas of the Cerrado and Pan- scientific information. tanal (outside of conservation areas) can help us a great deal to understand the dynamics of these Several characteristics of the physiognomy of the Cer- ecosystems. Today, horses, cows and feral hogs (all rado and Pantanal are anachronistic and reflect an introduced into the Pantanal less than 200 years ago), environment molded by the absence of large mammals, when in controlled densities, are important dispersers whether this is due to climatic changes or to the of large seeds and controllers of invasive herbs. anthropic effect of primitive man. The extensive consumption of plants by ants and termites (the More than two million nonnative animals, such as cattle, famous “cupinzeiros” of Emas), the reduced genetic buffaloes, horses and feral hogs wander the Pantanal variation of some plants, the low fertility of the soil, the (Mourão et al., 2003). We still know very little about clustered distribution of many vegetation species, and how this introduced megafauna molds the vegetation, the current importance of fire in the structuring of and if we should really completely remove this nonnative communities in the Cerrado, are some of the ecological fauna from the Pantanal. The impact of large herbivores patterns that should be looked at in terms of the will depend largely on their density within the absence of the pleistocenic megafauna. environment. In recent years, several researchers have been debating the impact of recent, and Pleistocene, Actually, the extinction of the megafauna did not stop to current ecosystems (Levin et al., 2002; during the Pleistocene/Holocene transition, and it is Ellsworth & McComb, 2003). This theme is fertile just one part of the history of man’s disturbances to ground ready to be tested in neotropical savannas . the planet. Today, many areas of the Cerrado, Atlantic Rainforest and the Amazon are suffering from a process However, where would this megafauna to be introduced of continual megafauna elimination. It is estimated that come from? Our circuses have about two thousand sixty million animals are hunted annually for con- animals for exhibition (almost always in humiliating sumption in the Amazon alone (Refdord, 1997). Add conditions). The great majority of these circuses are to this millions more that are captured in illegal live ani- succumbing financially, since the public increasingly mal trafficking. prefers circuses without animals. Additionally, there are animals in zoos that lack appropriate facilities to The first animals to disappear in areas impacted by maintain the animal’s well -being. hunting are, once again, the large mammals, such as woolly monkey (Lagothrix), muriqui (Brachyteles), Couldn’t we designate some areas in the Cerrado and tapirs and feral hogs. About 80% of the biomass of the Pantanal for experiments like those in the so-called the hunted animals is frugivorous-herbivore. In other “Pleistocene Parks?” A model of a Pleistocene Park words, its removal from the environment causes a already exists in Siberia, where researchers discovered that is reflected in the entire community. that the dominance of mosses in the tundras is a result Without frugivores, many plants are doomed to local of the removal of mammoths, horses and bisons, and extinction because of the loss of their dispersers (Peres that the reintroduction of the megafauna which survived & Roosmalen, 2002; Wright, 2003). hunting and climatic changes (such as bisons and horses), resulted in the appearance of a vegetation This reduction of fauna occurring in many ecosystems, dominated by grasses and bushes (Zimov et al., 1995). has caused several researchers to demonstrate that many fruit species with large seeds have their days The Pleistocene Parks would, obviously, not be placed numbered, because of an absence of dispersors. In in strictly protected areas. Certainly a lot of wetland some fragments of the Atlantic Rainforest the reduction farms and the Cerrado could house the “Pleistocene of fauna is so extreme that there are no seed dispersers Parks.” Obviously, the introduction of elephants, larger than a squirrel (Galetti et al., data not published). horses, guanacos and even hippopotami (similar to Therefore, the Pleistocene removal, as much as the Toxodons) is not an act of scientific imprudence, but current removal of megafauna, caused, and still cau- should be controlled and constantly monitored. The ses, a large impact on the vegetation, which is reflected accidental introduction of diseases is the largest threat in the entire ecosystem and directly affects the quality posed by programs of reintroduction, for this reason of life of human beings. the introduced animals should go through a rigorous health inspection and they should be constantly I believe that the plans to manage the Cerrado and the evaluated. Pantanal should take into account an historical process

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